Articles

How Many Species Are There on Earth?

RoBERT M. MAY

most notable problem being that some studies identify individual

This article surveys current answers to the factual ques-
tion posed in the title and reviews the kinds of informa-
tion that are needed to make these answers more precise,
Various factors affecting diversity are also reviewed.
These include the structure of food webs, the relative
abundance of species, the number of species and of
individuals in different cate~ries of body size, along with
other determinants of the commonness and rarity of
organisms.
species ("blue jays") whereas others deal with aggregates ("spiders,"
"copepods," or even "zooplankton"); some studies articulate indi-
vidual species of predators at upper levels but aggregate coarsely at
lower ttophic levels (3). Even so, some remarkable regularities
emerge from Cohen and Briand's analysis of these data (1, 2).
For one thing, the average number of other species with which
any one species interacts directly is consistently around 3 to 5 (4).
The number is consistently higher (average, 4.6) in relatively
constant environments than in fluctuating ones (average, 3.2).
There are also consistent and quantitative patterns in the propor-
tions of basal, intermediate, .and top predator species (those whose

0
VBR A Cl!NTURY AGO, DARWIN AND OTHERS PROVIDED
the broad outline of an answer to the question of how life
has evolved on Earth and how species originate. The next
question would seem to be how we use this basic understanding to
estimate-from first principles-how many species are likely to be
found in a given region or, indeed, on Earth as a whole.
Surprisingly, this question of "how many species?" has received
relatively little systematic attention, from Darwin's time to our own.
At the purely factual level, we do not know to within an order of
magnitude how many species of plants and animals we share the
globe with: fewer ):han 2 million are currently classified, and
estimates of the total number range from under 5 million to more
than 50 million. At the theoretical level, things are even worse: we
cannot explain from first principles why the global total is of the
general order of l07 rather than 104 or 10 10 .
This article first surveys various kinds of empirical and theoretical
links reach only upward, both ways, and only downward, respective-
ly); the ratios are 0.19 : 0.53 : 0.29, respectively (5). A similar
pattern of "link-scaling'' invariance is found for the ratio of links
among the four categories of basal-intermediate, basal-top, interme-
diate-intermediate, and intermediate-top (0.27: 0.08: 0.30: 0.35,
respectively). Most interestingly, these two quantitative patterns in
the proportions of species in different ttophic categories, and in
"link-scaling," can be deduced from the empirical observation that
each species is directly connected to roughly four others, along with
8.0
1.&
-------
7.0
6.0 /
/
/

studies that are helping to give us a better idea of how many species, I
/

or how many individual organisms, we might expect to find in a 5.0 I

given environment. Such studies include the sttucture of food webs,

patterns in the relative abundance of species, patterns in the number 4.0
of species or number of individuals in different categories of physical 0
size, and general observations about trends in the commonness or • 0
rarity of organisms. The article then reviews current evidence about
the total number of species on Earth, iridicating lines of research that • Birds
o Moths
could sharpen the estimates. We do not end up with a list of 2.0 a Gastropods
answers, but rather with a list of more sharply focused questions. • Plants
• Diatoms

The Structure of Food Webs 0 200 400 600 BOO

Cohen and Briand ( 1, 2) have compiled and analyzed a catalog
that now includes 113 food webs, embracing a wide variety of
natural environments (55 food webs from continental settings-23
terrestrial and 32 aquatic-along with 45 coastal and 13 oceanic
webs). The data for these food webs are of uneven quality, with the
This article was written while the author was in the Department of Biology, Princeton
University, Princeton, NJ 08544. His present address is Department of Zoology,
Oxford University, Oxford, OXl 3PS, England.
s
Fig. 1. A plot of S, the number of species, versus cr, the stand<!rd devi<~tion of
the log<~rithms of the rel<1tive <1bundances, for v<~rious communities of birds,
moths, g<~Stropods, plants, and diatoms. The d<~Shed line labeled 'I = 1.0
shows the relation between Sand cr for Preston's (15) "canonical" lognormal
distribution; the lines labeled 'I = 0.2 and 'I = 1.8 are the bounds to the
range of S-cr relations that might be expected from general mathematical
properties of the lognormal distribution, for large S and reasonable ranges of
values for the total number of individuals, N. The solid line is the mean
relation predicted by Sugihara's (17) model, and the error bars represent ±2
standard deviations about this mean. ·

I6 SEPTEMBER I988 ARTICLES I44I

Table 1. The distribution of 160 plant species from the Biological Flora of the
British Isles, classified into eight categories according to geographic distribu-
tion (wide or narrow), habitat specificity (broad or restricted), and local
abundance (somewhere large or everywhere small) (34).
Geographic distribution
Local
Wide Narrow
population
habitat specificity habitat specificity
size
Broad Restricted Broad Restricted
Somewhere large 58 7l 6 14
Everywhere small 2 6 0 3
the asswnption that the species are ordered in a cascade or hierarchy,
such that a given species can prey on only those below it and can be
preyed on only by those species above it in the hierarchy [an
asswnption that several authors (2, 6) have independently suggested
may follow from body-size considerations between predators and
their prey].
Other patterns in the ratios between nwnbers of interacting
species in different trophic levels are the subject of continuing
investigation. Hawkins and Lawton (7) have observed that food
chains comprising green plants, insect herbivores, and insect para-
sitoids include over half of all known species of metazoans, so that
understanding what determines the richness of parasitoid species
could be a major step toward understanding the diversity of
terrestrial communities. They analyzed data for 285 species of
herbivorous insects, from 42 families, in Britain, and found the
typical such species to be attacked by 5 to 10 species of parasitoids;
the number depends significantly on the geographical range of the
host insect, on the architecture of the host plant, and to a lesser
extent on a variety of other factors (7). Preliminary data suggest that
the tropics are roughly similar to Britain, in that herbivorous insects
are hosts to around five to ten species of parasitoids (8). Other
studies docwnent systematic patterns in the nwnber of phytopha-
gous insect species associated with different plant hosts (9) and in
the ratios between nwnbers of species of prey and predators of
various kinds ( 10).
It could be that many of these apparent patterns tell us more
about the workings of the hwnan mind, and about how we tend to
collect and categorize data, than they do about the natural world
( 11). Moreover, the populations in real food webs can have extreme-
ly complex dynamical behavior, with nonlinearities in density-
dependent factors producing cyclic or chaotic changes in abundance
and with unpredictable environmental fluctuations adding further
complications; it seems unlikely that the salient features of such
dynamical systems can be captured in static analyses of food web
graphs (12). These caveats and complications notwithstanding, the
patterns discussed above are intriguing. If they stand up to further
study, they could simplify the task of understanding diversity. It
could be, for example, that one need only understand what deter-
mines the nwnber of plant species, and then the total faunal diversity
could be deduced from appropriate rules.
Relative Abundance of Species
Real understanding offood webs in particular, and of diversity in
general, must go beyond the mere presence or absence of species to
an understanding of relative abundance. In early successional com-
munities, and in environments disturbed by toxins or "enriched" by
pollution, steeply graded distributions of species relative abundance
(SRA) are commonly seen, with a handful of dominant species
accounting for most of the individuals present. Conversely, in
relatively undisturbed "climax" communities consisting of many
species, relatively even distributions of relative abundance are typi-
cal; very often, such SRAs are distributed according to a "canonical
lognormal" distribution, as illustrated in Fig. l. Such trends in
SRAs show up in studies of old field succession (13). The effects of
pollution or other systematic disturbances reveal the same trend,
except that time effectively runs backward, so that the progression is
from evenness to dominance (14).
It is not surprising that the relative abundances within a fairly
large and relatively undisturbed group of species will be disturbed
lognormally. The relative abundances are likely to be governed by
the interplay of many more or less independent factors. It is in the
nature of the equations of population dynamics that these several
factors should compound multiplicatively, and the statistical central
limit theorem applied to such a product of factors implies a
lognormal distribution. This general observation, however, tells us
nothing about the relation between cr (the standard deviation of the
logarithms of the relative abundances) and S (the total nwnber of
species present). The puzzling fact is that very many assemblies have
SRAs that obey the canonical lognormal distribution, that is, that
have the unique relation between cr and S illustrated by the curve
labeled 'Y = l.O in Fig. l, although this curve represents just one of
an infinite family of possible lognormal distributions (15).
It has been conjectured that the canonical property may be merely
an approximate mathematical property of all lognormal distribu-
tions for large S; the dashed curves in Fig. l labeled 'Y = 1.8 and
'Y = 0.2 represent plausible boundaries to the cr-S relation on this
basis (16). The data put together by Sugihara (17) in Fig. l make it
clear, however, that real SRAs obey the canonical relation more
closely than can be explained by these mathematical generalities
alone. Sugihara has also suggested a biological mechanism that will
produce the observed patterns. He imagines the multidimensional
"niche space" of the community as being a hypervolwne broken up
sequentially by the component species (with any fragment being
equally likely to be chosen for the next breakage, regardless of size),
such that each of the S fragments denotes the relative abundance of a
species. Although the biological status of this asswnption is debat-
able, it generates patterns of SRA in accord with a large nwnber of
data (the solid line in Fig. l shows the mean relation between Sand
103
"'
cu
'()
cu
a.
."'
0 102
cu
.c
E
::1
z
10
,...-----
_____ _____ ,
0 2 3 4 56 7
log 10[Weight (g)]
Fig. 2. The numbers of species, S, of all terrestrial mammals (solid
histogram) and of British mammals (dashed histogram), excluding bats, are
shown distributed according to mass categories (mass expressed in grams)
(18, 19). Note the doubly logarithmic scale. The thin dashed line illustrates
the shape of the relationS- L - 2 , where Lis the characteristic length (20).
SCIENCE, VOL. 241
o- predicted by the model, and the error bars show the range of ±2 however, that the structure of the habitat-and hence the number of
standard deviations about the mean). Such a fit does not, of course, possible ways of making a living-is unlikely to scale linearly with L
validate the model; it is possible that other biological assumptions (26). Consider, fur example, the circumference of a large tree, or any
could produce similar distributions of SRA. other "one-dimensional" object. If we measure it on a 10-cm scale,
One problem with essentially all the data that have been compiled we get one answer. On a 1-cm scale, we will often get another, larger
for SRA is that they focus on particular taxonomic groups ("birds," answer. A yet larger answer would be obtained on a 1-mm scale, and
"moths"). To understand how communities are assembled, it may so on. The circumference of the tree is .thus not simply one
be more relevant to inquire about the relative abundance within dimensional but has a "fractal dimension," D, such that the per-
ecologically similar groups (putting birds together with bats and ceived length, e(A.), deEepds on the step-length of measurement, A.,
some large insects, for instance). as x_l-D: e(A.) = cA. 1- , where c is a constant. If D = 1.5, for
example, a 10-fold reduction in the measurement scale (from, say lO
em to 1 em) will result in the apparent length increasing by a factor
10°·5 = 3. Morse et al. applied these notions to measure the profiles
Number of Species Versus Physical Size of various kinds of vegetation at different scales, concluding that D
A variety of other patterns in the distribution and abundance of for such habitats ranged from around 1.3 to around 1.8, with an
organisms have received little attention. For example, how many average around 1.5 (Fig. 3). That is, for herbivorous insects that
species do we expect to find in different categories of physical size, exploit their surroundings in an essentially one-dimensional way
within a given region? (using the edges ofleaves, or the like) a 10-fold decrease in physical
The meager amount of available information bearing on this size produces a roughly 3-fold increase in the apparently available
question is reviewed elsewhere (18). Figure 2 gives one repre-
sentative study, showing the way in which all 3000 or so mammali- Table .2. The number of species (to within an order of magnitude) in the
different animal phyla, classified according to the habitat of adult animals.
an species, excluding bats and marine mammals, are apportioned Most phyla are predominantly marine and benthic, some exclusively so. The
among mass classes (19). A corresponding analysis, but restricted to numbers 1 through 5 indicate the approximate number of recorded living
the mammal species of Britain, again excluding bats and marine species: 1 means I to I02 ; 2 means I02 to 103; 3 means 103 to 104 ; 4 means
mammals, is also shown in Fig. 2 (18). Although Britain's mammals 104 to 105 ; and 5 means 105 or more. After (38). Abbreviations: B, benthic;
P, pelagic; M, moist; X, xeric; Ec, ecto; and En, endo.
appear to obey the global pattern of species versus size, appropriate-
ly scaled down, this may not be true in general; there is no a priori Habitat
reason to expect the species-size patterns for faunal assemblies from Phylum
relatively small areas to be the same as those from large (and Marine Freshwater Terrestrial Symbiotic
Subphylum
correspondingly more environmentally diverse) areas. B p B p M X Ec En
Figure 2 and similar analyses represent rough assessments of the
Porifera 3 1 I
facts. Very few ideas have been advanced in explanation of these Placowa 1
facts about species-size distributions. Hutchinson and MacArthur Orthonectida 1
(20) have advanced arguments for expecting an L - 2 relation be- Dicyemida I
tween the number of species and the characteristic length of Cnidaria 3 2 I 1 1
constituent individuals, L. The argument is essentially that, for Ctenophora 1 I
Platyhelminthes 3 1 3 2 1 4
terrestrial organisms, the world is seen as two-dimensional, and Gnathostomulida 2
therefore the possibility of finding new roles (and thence new Nemertea 2 1 1 1 1
species) may scale as L - 2 • This conjectured L - 2, or M- 213 , relation, Nematoda 3 1 3 1 3 I 3 3
where M is mass, is illustrated by the dashed straight line in Fig. 2. Nematomorpha 2
Acanthocephala 2
Rotifera 1 1 2 2 1 I 1
Gastrotricha 2 2
Number of Individuals Versus Physical Size Kinorhyncha
Loricifera
i
1
Other patterns can be sought in the relation between numbers of Tardigrada I 2 I
individuals and their physical size (mass or characteristic length). Priapula 1
For example, in a particular region, how is the number of individual Mollusca 5 1 3 3 I 1 1
Kamptozoa 1 1 I
animals in the size class from 0.1 to 1 em related to the number in Pogonophora 2
the class from 1 to 10 em (21, 22)? SipUncula 2 1
In particular, Morse et al. (21) and Brown and Maurer (23) have Echiura. 2
collated data about populations of phytophagous insects and of Annelida 4 1 2 3 2
Onychophora 1
birds, respectively, and have advanced qualitative explanations for Arthropoda
these data. Morse et al. began with the assumption that roughly Crustacea 4 3 3 2 2 2 2
equal amounts of energy flow through each size category; although Chelicerata 2 I 2 2 4 3 2 1
very unlikely to be true in general, this assumption is supported by Uniramia I I 3 2 5 3 2 2
some evidence from organisms ranging widely in size (24). Given Chaetognatha I I
this assumption, along with the usual manner in which metabolic Phoronida I
Brachiopoda 2
costs become relatively larger at smaller sizes, the total number of Bryowa 3 I
individuals, N, in the size class with characteristic mass M and length Echinodermata 3 I
L may be expected to scale as N- M-0 ·75 - L - 2·25 (25). That is, for Hemichordata I
a 10-fold decrease in characteristic length we would, on this basis, Chordata
Urochordata 3 I
expect a roughly 180-fold increase in the total number of individ- Cephalochordata I
uals. Vertebrata 3 3 2 3 3 3 I I
Recent insights into the fractal geometry of nature suggest,
16 SEPTEMBER 1988 ARTICLES 1#3
habitat; for creatures that exploit their environment in an essentially 10% of the globe, over 1%, and so on? Hanski (30), Brown (31),
two-dimensional way (using surfaces rather than edges), the effect Root (32), Rapoport (33), and others have made a start toward
must be squared, so that a 10-fold decrease in physical size produces answering this question, for diverse collections of organisms includ-
an effectively 10-fold increase in apparent habitat. These two factors ing vascular plants, intertidal invertebrates, terrestrial arthropods,
(the one-dimensional factor 3 and the two-dimensional factor 10) planktonic crustaceans, and terrestrial vertebrates (especially birds),
are likely to bound the range of possibilities found in actual but much remains to be learned.
assemblies of insects. Intuitive ideas about commonness and rarity usually make refer-
Combining these fractal aspects of habitat perception with the ence both to geographical distribution and to local abundance. Such
metabolic considerations discussed above, Morse et al. concluded considerations often swirl together in ways that make it difficult to
that a 10-fold decrease in characteristic length, L, is likely to produce define exactly what constitutes a rare species. One type of rareness is,
an increase in N that lies between 500 and 2000 (that is, roughly for example, exhibited by the silver sword, Argyroxyphium macro-
between 3 and 10 times 180). As shown in Fig. 4, this very rough cephalum, that grows only in the crater of the Haleakala volcano on
expectation is borne out surprisingly well by data for the number of Maui. Although there are around 50,000 individuals in the large,
individual arthropods of different body lengths found on vegetation
in places ranging from primary forests, primary riparian vegetation, Table 3. A rough indication of the relative effort devoted to animals from
and secondary vegetation in the New World Tropics to temperate different taxonomic groups is given by the average number of papers listed in
habits, for example, birch trees on Skipwith Common in Yorkshire. the Zoological Record, 1978 through 1987 (54).
The study by Morse et al. is a frankly speculative one. I have Average
chosen to highlight it because it provides an explicit example where number of Papers
Phylum Approximate
our thinking about aspects of population abundance and diversity Subphylum publications
number of per
needs to acknowledge that nature is often not Euclidean but rather Class per year species
recorded
(coefficient per
may have fractal geometry, with organisms existing in spatial and Order
of variation,
species
year
temporal frameworks that are, as it were, jagged on every scale (27). in percent)
This is an example where new mathematical concepts interact with
biological ideas in potentially surprising ways (the chaotic behavior Protowa 3,900 (10) 260,000 0.15
Porifera 190 (22) 10,000 0.02
of many simple population models is another example). Coelenterata 740 (12) 10,000 0.07
Echinoderma 710 (15) 6,000 0.12
Nematoda 1,900 ( 1) 1,000,000 (?) 0.002
Annelida 840 ( 9) 15,000 0.06
Species Numbers, Species Abundance, and Brachiopoda 220 (14) 350 0.63
Body Length Bryowa 160 (15) 4,000 0.04
Entoproctra 7 (53) 150 0.04
Still other studies have focused on empirical relations between the Mollusca 1,000 ( 8) 100,000 0.04
abundance of individual species and the body size of constituent Arthropoda
Crustacea 3,300 ( 9) 39,000 0.09
individuals (25, 28). I think any eventual understanding of the total
Chclicerata
number of species in a given environment, and thence ultimately of Arachnida 2,000 ( 6) 63,000 0.03
the diversity of life on Earth, will need to be based on a clear Uniramia
understanding of the interplay among all the factors discussed Insecta 17,000 7) 1,000,000 (?) 0.02
above. Yet most of the few existing studies have singled out one or Coleoptera 2,900 6) 300,000 0.01
Diptera 3,200 7) 85,000 0.04
other aspect (species size, species abundance) from the interwoven Lepidoptera 3,500 9) 110,000 0.03
mosalC. Hymenoptera 2,200 9) 110,000 0.02
Exceptions are the work on birds by Brown and Maurer (23) and Hemiptera 1,700 7) 40,000 0.04
the recent study by Morse et al. of the relations among species Chordata
Vertebrata
number, species abundance, and body length for 859 species of Pisces 7,000 (13) 19,000 0.37
arboreal beetles in lowland rain-forest trees in Borneo (29). Figure Amphibia 1,300 (12) 2,800 0.47
SA summarizes the results, showing the total number of species in Reptilia 2,400 ( 7) 6,000 0.41
diiferent categories of population abundance and physical size (both Aves 9,000 (10) 9,000 1.00
plotted logarithmically); Fig. 5, B through E, correspondingly Mammalia 8,100 (12) 4,500 1.80
shows the number of species in different trophic categories. Al- Mammalian orders
Monotremata 20 3 6.8
though Fig. 5 does have some interesting structural details [for Marsupialia 269 266 l.O
discussion, see (29) ], it is essentially simple. It is encouraging that Insectivora 270 345 0.8
Fig. 5 has the basic features one would have guessed from the Dermoptera 2.2 2 l.l
separate studies of species abundance, species size, and abundance Chiroptera 402 951 0.4
Primates 956 181 5.3
size in different groups, as discussed above. Edemata 38 29 1.3
Pholidota 5 7 0.7
Lagomorpha 173 58 3.0
Rodentia 1,538 1,702 0.9
Commonness and Rarity Cetacea 360 76 4.8
Carnivora 1,157 231 5.0
In the discussion above, some of the species found in a given Tubulidentata 2.7 1 2.7
region are confined to tlut region, whereas others (which are part of Proboscidea 94 2 47
the species-size and other distributions in the region) are distributed Hyracoidea l2 11 l.O
much more widely. Partly for this reason, and partly for its intrinsic Sirenia 43 4 10.8
Perissodacryla 142 16 8.9
interest, it would be nice to know more about the distribution of Artiodactyla 187 6.0
1,124
geographical ranges within different taxonomic groups of species. Pinnipcdia 218 33 6.6
What fraction of all bird species, for example, range globally over
1444 SCIENCE, VOL. 241
local population of this plant, its restriction to the one volcanic
crater would make it very rare by most definitions. Another type of
rareness is exhibited by the grass Setaria geniculata, which is found
from Massachusetts to California and on down through tropical
South America to Argentina and Chile but which is not abundant
anywhere. This grass is rare in the sense that its populations are
"chronically sparse" (34) everywhere in its broad range.
There have been a variety of proposals for codifYing ideas about
commonness and rarity. In particular, Rabinowitz et al. (34) have
considered three different kinds of questions that arise in thinking
about rarity: (i) is the species distributed over a broad geographical
area, or is it endemic to some restricted location; (ii) whatever its
range, is the species found in a wide variety of habitats, or is it
specialized to one kind of site; and (iii) is the species abundant
somewhere in its range, or are its numbers everywhere small. These
three considerations combine to give eight categories, only one of
which (broad distribution, unspecialized habitat, large populations)
ordinarily corresponds to the species being called "common."
Rabinowitz et al. noted that the archetypal "rare" species, with
narrow distribution, specialized habitat, and small numbers, repre-
sents only one of several different kinds of rarity. These investigators
pursued their ideas by applying them to the plants surveyed in the
Biological Flora of the British Isles (which gives detailed distribution
maps and notes about the habitat and population of 177 of the 1822
native British plants). Rabinowitz et al. asked 15 colleagues to
classify each of the 177 species according to the eightfold category
scheme described above (35). This process gave clear consensus for
160 of the 177 species, and the results are summarized in Table l.
Most species ( 149 versus ll) are abundant somewhere, and most
species (137versus 23) have a wide geographical range (Table 1). A
narrower majority (94 versus 66) have restricted habitat specificity.
Of the eight categories, species with wide ranges and large
population sizes, but restricted habitat specificities, predominate (71
1oo,ooo B
10,000
A
1,000
0=1.30
(high
magnification)
100
10
0=1.55
(low magnification)
1+-----r---~----~
1 10 100 1,000
No. of squares on one
side of grid
Fig. 3. (A) Photographs of plants at various magnifications were placed
under a grid by Morse et al. (21). The number of squares entered by the
outline of the plant were counted, starting with a coarse grid of two large
squares on one side, then 2n squares, with n varying from 2 to 6 or 7,
depending on the grid size. For ease of representation, the plant's leaves in
this figure are drawn flat; in reality they are oriented at all angles with respect
to the grid. Also for clarity, the progressively finer divisions are only
illustrated in one comer of the figure. The logarithm of the number of
squares entered by the outline of the plant is then plotted against the
logarithm of the number of squares along one side of the grid, as shown in
(B). The slope of the line equals the fractal dimension, D. (B) Data gathered
in this way for Virginia creeper, photographed without leaves in early spring.
The twigs were photographed at one scale, then parts of the same twigs were
rephotographed at a higher magnification, permitting D to be estimated at
two levels of resolution.
16 SEPTEMBER 1988
Table 4. Estimated numbers of host-specific canopy beetles on Luehea
seemannii, classified into trophic groups (49).
Estimated Estimated
Trophic Number of fraction number of
group species host-specific host-specific
(%) species
Herbivores 682 20 140
Predators 296 5 15
Fungivores 69 10 7
Scavengers 96 5 5
Total 1100+ 160
of 160 species, or 44% ). Most of these "rare habitat" species are
specialists of marsh, sand dunes, bogs, or forest floors; wherever
their habitat exists, they are predictably present (36). The category
that is conventionally called "common" comes a close second,
exemplified by species such as heather, Calluna vulgaris, or English
oak, Quercus robur (58 of 160 species, or 36%). The remaining six
categories are all less well represented, collectively accounting for
only 20% of the total. The most frequent of these six are what are
usually called "endemic rarities," specializing in one type of habitat
but abundant in that habitat (14 of 160 species, or 9%); the Lady
Orchid, Orchis purpurea, in Kent is an example. Other categories are
uncommon, and one is unrepresented in the British flora: Rabino-
witz et al. found no species with small populations in a variety of
habitats but with a narrow geographic distribution. The absence of
this category may reflect the small sample size, or it may reflect
ecological mechanisms that are not yet fully understood.
We need more of these kinds of empirical studies of the multifac-
torial determinants of commonness and rarity (37). Not only do
such studies illuminate fundamental questions about diversity, but
they have practical implications for conservation biology. For
instance, Table 1 helps justify the attention traditionally given by
conservationists to "endemic rarities": not only are these species,
with their narrow ranges and restricted habitat specificities, easily
destroyed, but they are also numerically the most prevalent category
of rare plants. Rabinowitz et al. speculated, moreover, that a better
understanding of how endangered and extinct species are appor-
tioned among their eight categories (or among other, alternative
categories) may offer "clues about the causes of the endangered
state" (34, p. 200).
How Many Living Species Have Been
Recorded?
So far, this article has dealt with issues that must be resolved if we
are ever to estimate the number of species in a given region, or on
Earth, from basic principles. The second part of the article now
reviews our current ignorance about the simple facts of how many
species there actually are.
Living things may be divided into five kingdoms, distinguished
by different levels of cellular organization and modes of nutrition.
Two of these kingdoms, the prokaryotic monerans and the eukary-
otic protists, comprise microscopic unicellular organisms, and to-
gether they account for something like 5% of recorded living
species. The fungal and plant kingdoms represent roughly another
22% of species. The animal kingdom thus comprises the majority
(more than 70%) of all recorded living species (38). Table 2 gives a
rough account of how the species in the different animal phyla are
apportioned according to the habitat of the adult creatures; each
phylum represents a distinct body plan, with fundamental differ-
ences that distinguish it from all the others (39).
ARTICLES 1445
 10,000
\ A \ 8 \ c \ D \ E
\ \ \ \ \
\ \ \ \ \
\
0 •
\ \ \ \

.. ..
1,000 \ \ \

....
\ 0 \

..
~ \ \ \
\
• 0
\
tA
\ \ \ \ \ \ \ \

. '\\
iU \

'\ '
\
'' •. '
\
0
:I 0
\ \ \ \

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> 100 \ \ \
": \ \

' ... '' ''

\
0 • 0
'5 \ \ \
.5 \ \ \
''
\ \
'' \ \
0

'' .. . '
• 00

''
0 \ \ \ •'I> \

' '
0 0.
'! \
\ \
,.. ....
d \ \ \
z
... . '
0 00

.. . '
'
' .- '
10 \ \ II • \ \ \ \ \ \

''
00
\ \ : \
\ \ \ \ \ • 0

\
'\ \ \ \ \ \ \

'
'
0 ••

\• \
0 •
\ \ \ \
1 10 100 10 100
1 10 100 10 100
Body length, L (mm)
Fig. 4. Data plotted by Morse et a/. (21) on the number of individual Skipwith Common, North Yorkshire. The lower bound prediction that, for
arthropods (mainly insects) of different body lengths, collected from vegeta- an order of magnitude decrease in body length, there should be a roughly
cion: (A) understory foliage in primary forest in Costa Rica; (B) Osa 500-fuld increase in the number of individuals, is indicated by the lower
secondary vegetation (solid dots) and Kansas secondary vegetation (open dashed line on each graph. The upper bound prediction-roughly 2000-fold
dots); (C) Tabago primary riparian vegetation (solid dots) and Icacos increase for an order of magnitude decrease in body length-is shown by the
vegetation (open dots); (D) understory foliage in cacao plantations in upper dashed line.
Dominica (solid dots) and in Costa Rica (open dots); and (E) birch trees at

Table 2 shows that most phyla are fmmd in the sea, and more genetic distance between mammals and fishes. Relatively easy
particularly in benthic environments; many phyla are found only in exchange of genetic material among different "species" of microor-
benthic habitats. On the other hand, by far the most abundant ganisms could mean that basic notions about what constitutes a
category of recorded living species is terrestrial insects. To a rough species are necessarily different for vertebrates than for bacteria. But
approximation and setting aside vertebrate chauvinism, it can be I think there are likely also to be systematic trends toward greater
said that essentially all organisms are insects. Hutchinson (40, p. lumping of species of small and relatively less-studied organisms,
149) has suggested that "the extraordinary diversity of the terrestrial and toward greater splitting as we approach the furries and feather-
fauna, which is much greater than that of the marine fauna, is clearly ies.
due to the diversity provided by terrestrial plants." Although it is In Table 3, I attempt to give a rough impression of how the
true that in the sea vegetation does not form a structured environ- efforts of professional taxonomists and systematists are currently
ment (except close to shore) and that species generally have large distributed among the major groups of organisms. Obviously the
geographical ranges (and the oceans are contiguous), closer exami- vertebrates, which comprise only 3% of all animal species, receive a
nation suggests that there are subtle boundaries to dispersal in the disproportionate amount of attention. One result is that new birds
sea and that latitudinal zonation is often more marked in the sea continue to be found at the rate of about three species per year
than on land (41). Viewing these questions in another light, Ray (against a total of around 8000 species), and new mammals at the
(42) has observed that although the sea contains only 20% of all rate of around one genus per year (against a total of around 600
animal species, it contains systematically higher proportions of genera), which contrasts with the possibility that there may be more
higher taxonomic units, culminating in 90% or more of all classes or than ten insect species for every one yet classified (45).
phyla (largely because all phyla are found in the sea, and the bulk of Setting all these reservations and biases aside, the total number of
classes are exclusively marine). These facts make it plain that the living organisms that have received Latin binomial names is current-
factors influencing how many species there are in any one place- ly around 1.5 million or so (46). Amazingly, there is as yet no
food web structure, relative abundance, species-size patterns, and so centralized computer index of these recorded species. It says a lot
on-<:an operate differently in different environments and on differ- about intellectual fashions, and about our values, that we have a
ent spatial scales. computerized catalog entry, along with many details, for each of
Any interpretation of information about diversity, such as that several million books in the Library of Congress but no such catalog
summarized in Table 2, is clouded by uncertainties about how for the living species we share our world with. Such a catalog, with
different two groups of organisms have to be before we call them appropriately coded information about the habitat, geographical
different species, and by the fact that some taxa (for example, distribution, and characteristic abundance of the species in question
vertebrates) have been studied in vastly more detail than others (for (no matter how rough or impressionistic), would cost orders of
example, mites). Even within very well studied groups, some magPitude less money than sequencing the human genome; I do not
workers recognize many more species than others. This is especially believe such a project is orders of magnitude less important.
the case for organisms that can reproduce asexually; thus some Without such a factual catalog, it is hard to unravel the patterns and
taxonomists see around 200 species of the parthenogenetic British processes that determine the biotic diversity of our planet.
blackberry, others see only around 20 (and a "lumping" invertebrate
taxonomist may concede only 2 or 3). Some strongly inbreeding
populations are almost as bad, with "splitters" seeing an order of
magnitude more species than do "lumpers" (43). At a more funda-
How Many Living Species Are There?
mental level, Selander (44) observed that different strains of what is Until recently, the total number of species was thought to be
currently classified as a single bacterial species, Legionella pneumo- around 3 million to 5 million. This estimate was obtained roughly as
phila, have nucleotide sequence homologies (as revealed by DNA follows (46). For the species of mammals, birds, and other larger
hybridization) of less than 50%; this is as large as the characteristic animals that are relatively well enumerated, there are roughly twice
14+6 SCIENCE, VOL. 241
as many species in tropical regions as in temperate ones. The total
number of species actually named and recorded is around 1.5
million, and two-thirds of these are found in temperate regions.
Most of these are insects. But most insects that have actually been
named and taxonomically classified are from temperate zones. Thus,
if the ratio of numbers of tropical to temperate species is the same
for insects as for mammals and birds, we may expect there to be
something like two yet-unnamed species of tropical insects for every
one named temperate species. Hence the overall crude estimate of a
total of roughly three times the number currently classified, or
around 3 million to 5 million.
This estimate is open to several questions. For one thing, the total
includes relatively few species of bacterial, protozoan, and helminth
parasites, largely because such parasites are usually studied in
connection with economically important animal hosts. But it could
be that essentially every animal species is host to at least one
specialized such parasitic species (47), which would immediately
double the estimated total. For another thing, the Acarina (mites),
both tropical and temperate, are even less well studied than tropical
insects; it was largely tropical insects that carried the estimate from
the known 1.5 million to 3 million to 5 million, and mites could
carry it significantly higher.
An indirect approach to the question of the number of species
whose body size is small is through studies of species-size relations,
such as that in Fig. 2. Figure 6 depicts a very crude estimate of the
global totals of terrestrial animal species in different size categories
(classified, on a logarithmic scale, according to characteristic body
A
Number of species
log Abundance § Cll log Body size
per species e ~
~ 105
-~~~
~~E~
~~
Fig. 5. (A) The height of each intersection is proportional to the number of
beede species that have a particular combination of body length [plotted
logarithmically on a scale that extends from 0.5 mm, "small," to 30 mm,
"large"] and abundance [plotted as logarithms to the base 2, on the
conventional "octave" scale of Preston (15)]; for details, see (29). (B through
E) The same information for the separate beede guilds of herbivores,
predators, fungivores, and scavengers, respectively (29).
16 SEPTEMBER 1988
length); the data in Fig. 6 are the result of a multitude of rough and
uncertain estimates (18). The dashed line indicates the scaling of
numbers of species as L -z (20); the fractal considerations reviewed
in connection with Figs. 3 and 4 suggest the scaling might more
appropriately be somewhere between L -t.s and L -J (48). Whatever
the detailed scaling relation at larger body sizes, it dearly breaks
down for organisms whose characteristic body length is significantly
below 1 em. But these are exactly the same creatures-insects, mites,
and the like-that have received relatively little attention from
taxonomists. Because we lack a fundamental understanding of the
size-species relation itself, there is no reason to expect a simple
extrapolation of the scaling law for large sizes to estimate accurately
the number of unclassified smaller species. It is, however, interesting
that the total number of species obtained by extrapolating down to
around 1 mm or so is in the range 10 million to 50 million.
A sounder basis for an upward revision of the estimated number
of species comes from Erwin's studies of the insect fauna in the
canopy of tropical trees (49). Using an insecticidal fog to "knock
down" the canopy insects, Erwin found that most tropical arthro-
pod species appear to live in the tree tops. This is not so surprising,
because this is where there is most sunshine as well as most green
leaves, fruits, and flowers.
Erwin's original studies (49) were on canopy-dwelling beetles
(including weevils) collected from Luehea seemannii trees in Panama
over three seasons. He found more than 1100 species of such
beetles, distributed among the categories of herbivore, predator,
fungivore, and scavenger as shown in Table 4. To use this informa-
tion as a basis for estimating the total number of insect species in the
tropics, one needs to know what fraction of the fauna are specific to
the particular tree species or genus under study; unfortunately, there
are essentially no data bearing on this point. Erwin estimated 20%
of the herbivorous beetles to be specific to Luehea (in the sense that
they must use this tree species in some way for successful reproduc-
tion) (Table 4); the overall answer is more sensitive to this guess
than to the corresponding figures of 5%, 10%, and 5% for predator,
fungivore, and scavenger beetles, respectively. In this way, one
arrives at the estimate of around 160 species of canopy beetles
specific to a typical tropical tree.
106
Ul 104
.9!
~
Q.
Ul
...
0 103
Cll
,g
E
:I
z
102
10
1
-0.5 0 0.5 1.5 2 2.5 3 3.5
log 10 [Length (mm)]
Fig. 6. A crude estimate of the distribution of number of species of all
terrestrial animals, categorized according to characteristic length L. The
dashed line indicates the relation S- L - 2 , as in Fig. 2 (S is number of
species) [after (18)]. The question mark emphasizes the crudity of these
estimates and the inadequacy of the data for small size classes.
ARTICLES 1#7
 Several other assumptions are needed to pyramid this estimate of
160 host-specific species of canopy beetles per tree to 30 million
species in total. Slightly simplified, the argument runs as follows.
First, Erwin noted that beetles represent 40% of all known arthro-
pod species, leading to an estimate of around 400 canopy arthropod
species per tree species. Next, Erwin suggested the canopy fauna is
at least twice as rich as the forest-floor fauna and is composed mainly
of different species; this increases the estimate to around 600
arthropod species that are specific to each species or genus-group of
tropical tree. Finally, using the estimate of 50,000 species of tropical
trees (50), Erwin arrived at the possibility that there are 30 million
tropical arthropods in total. This estimate has been widely cited,
often without full appreciation of the chain of argument underlying
it.
Although it is easy to cavil at each step in Erwin's argument, the
work is important in providing a new and focused approach to the
problem of estimating how many species there are. Erwin does not
so much answer the question as define an agenda of research.
First, the overall estimate depends almost linearly on the necessar-
ily arbitrary assumption that 20% of the herbivorous beetles are
found only on one species or genus-group of tree; changing this
number to 10% would halve the global estimate to 15 million
species. I think it likely that insects feeding in the canopies of rain-
forest trees could be significantly less specialized in their use of food
plants than are temperate insects, in order to help them deal with the
sparse distribution of many tropical trees. Experiments that
"knocked down" the canopy insect fauna from each of many
neighboring trees of different species could shed light on these issues
and provide a firmer basis for the estimates in the last column of
Table 4 (51).
Second, the fact that 40% of taxonomically classified arthropod
species are beetles is of doubtful relevance if, in truth, essentially all
arthropods are unclassified tropical canopy dwellers. What we need
to know is the fraction of the canopy fauna that are beetles. Again,
this information could be obtained by systematic studies of the
overall arthropod fauna in the canopies of a variety of tropical trees.
Third, the assumption that there are roughly two canopy species
for each forest-floor species is also amenable to systematic study.
Such studies should, in my view, reach below the forest floor into
the soil, attempting to get a better idea of the species diversity of
decomposing animals (including nematodes and other helminths)
and other soil-dwellers.
More generally, I believe our ignorance of tropical mites-to
name but one group--is at least as great as the ignorance about
beetles and other arthropods that Erwin has exposed. These other
groups may be similarly diverse. One proposal that is ambitious by
ecological standards (although not by those in the physical sciences)
is to assemble a team of taxonomists, with a comprehensive range of
expertise, and then make a rough list of all the species found in one
representative hectare in the tropical rain forest; it would be better
to census several such sites (52). Until this is done, I will not trust
any estimate of the global total of species.
Coda
For most of the history of life on Earth that is preserved in the
fossil record, rates of extinction and rates of speciation have been
roughly commensurate. If, however, we assume that something like
half the extant species evolved in the last 50 million to 100 million
years and that maybe half of all extant species will become extinct in
the next 50 to 100 years if current rates of tropical deforestation
continue, then contemporary rates of speciation are of order 1
million times slower than rates of extinction (53). Were speciation
I448
rates plotted as the y-axis on a graph 10 em high, then on the same
scale extinction rates would require an x-axis extending 100 km.
These circumstances give a special urgency to the kinds of studies
called for above. Unlike essentially all other scientific disciplines,
conservation biology is a science with a time limit, with the clock
ticking faster as the human population continues to increase. We
need to understand the world of living things for the same
fundamental reasons that we need to understand the physics of the
unimaginably small and of the unimaginably large. We also need
such understanding to manage the biosphere in a sustainable way
(which we do not appear to be doing at present) and to design
rational strategies for preserving some habitats while exploiting
others in ways that allow some fraction of the original flora and
fauna to persist. I believe future generations will find it blankly
incomprehensible that we are devoting so little money and effort to
the study of these questions.
REFERENCES AND NOTES
l. The details of 40 of these webs are fully docwnented in J. E. Cohen, l'ood Webs and
Niche Space (Princeton Univ. Press, Princeton, NJ, 1978) and F. Briand, Ecology
64, 253 (1983). The frequency distribution of chain lengths and other properties
have been reported for all 113 webs in J. E. Cohen, F. Briand, C. M. Newman,
Proc. R. Soc. London Ser. B 228, 317 (1986).
2. F. Briand and J, E. Cohen, Science 238, 956 (1987).
3. Other problems in the compilation of such catalogs of food webs include the
following: one wants the "community" web, not a subset of species traced up from
one resource or down from one top predator; one must decide, arbitrarily, which
links are regarded as too weak or too unusual to list; different researchers may have
followed different procedures or had different biases in tabulating individual webs;
and so on. For further discussion, see R. M. May, Nature 301, 566 (1983).
4. J, E. Cohen, C. M. Newman, F. E. Briand, Proc. R. Soc. London Ser. B 224, 449
(1985).
5. J. E. Cohen and C. M. Newman, ibid., p. 421. This study is based on 62 food webs
only.
6. P. H. Warren and J. H. Lawton, Oecologia (Berlin) 74, 231 (1987).
7. B. A. Hawkins and J. H. Lawton, Nature 326, 788 (1987).
8. J. H. Lawton, personal communication.
9. These patterns depend significandy on the range of the host plant: D. R. Strong, J.
H. Lawton, T. R. E. Southwood, Insects on Plants (Blackwell, Oxford, 1984); H. C.
Godfray,]. Econ. Entomol. 9, 163 (1984); P. W. Price, Evolutionary Biology of
Parasites (Princeton Univ. Press, Princeton, NJ, 1980).
10. M. J. Jeffries and J. H. Lawton, Freshwater Bioi. 15, 105 (1985); F. Briand and}. E.
Cohen, Nature 307, 264 (1984).
11. This nwnber of direct interactions with 3 to 5 other species is disconcertingly
evocative of Miller's observation d1at people tend to recall 7 ± 2 items for lists of
very varied lengths [G. A. Miller, Psychoanal. Rev. 63, 81 (1956)].
12. R. T. Paine,]. Anim. Ecol. 49, 667 (1980).
13. F. A. Bazzaz, Ecology 56, 485 (1975).
14. D. Tilman, Resource CompetitioH and Community Structure (Princeton Univ. Press,
Princeton, NJ, 1982); R. M. May, Ed., Theoretical Ecology: Principles and Applications
(Sinauer, Sunderland, MA, 1981).
15. F. W. Preston, Ecology 43, 185 (1962); R. H. MacArthur and E. 0. Wilson, Island
Biogeography (Princeton Univ. Press, Princeton, NJ, 1967).
16. R. M. May, in Ecology of Species and Communities, M. Cody and J. M. Diamond, Eds.
(Harvard Univ. Press, Cambridge, MA, 1975), pp. 81-120.
17. G. Sugihara, Am. Nat. 116, 770 (1980).
18. R. M. May, in Diversity of Insect Faunas, L. A. Mound and N. Waloff, Eds.
(Blackwell, Oxford, 1978), pp. 188-204.
19. L. Van Valen, Evolution 27,27 (1973). The pattern in Fig. 2 (and in Fig. 6) could
be somewhat changed if the large mammals that have become extinct over the last
million years or so are included.
20. G. E. Hutchinson and R. H. MacArthur, Am. Nat. 93, 117 (1959).
21. D. R. Morse, J. H. Lawton, M. M. Dodson, M. H. Williamson, Nature 314,731
(1985).
22. D. Griffiths, Am. Nat. 127, 140 (1986); D. Strayer, Oecologia (Berlin) 69, 513
(1986).
23. J. H. Brown and B. A. Maurer, Am. Nat. 130, 1 (1987).
24. E. D. Odwn, Fundamentals of Ecology (Saunders, Philadelphia, 1953).
25. R. H. Peters, The Ecological Implications of Body Size (Cambridge Univ. Press,
Cambridge, MA, 1983).
26. B. B. Mandelbrot, Fractals: Form, Chance, and Dimension (Freeman, San Francisco,
1977); C. Loehle, Spec. Sci. Tech. 6, 131 (1983).
27. R. M. May, in Ecological Concepts, J. M. Cherrett, Ed. (Blackwell, Oxford, in press).
28. J. Damuth, Nature 230, 699 (1981); R. H. Peters and J. V. Raelson, Am. Nat.
124, 498 (1984); R. H. Peters and K. Wassenberg, Oecologia (Berlin) 60, 89
(1983); J. H. Brown and B. A. Maurer, Natnre 324, 248 (1986).
29. D. R. Morse, N. E. Stork, J. H. Lawton,]. Ecol. Entomol. 13, 25 (1988).
30. I. Hanski, Oikos 38, 210 (1982).
31. J. H. Brown, Am. Nat. 124, 255 (1984).
32. T. Root, Atlas of Wintering North American Birds (Univ. of Chicago Press, Chicago,
SCIENCE, VOL. 24-I
 1988).
33. E. H. Rapoport, Areogeography: Geographical Strategies of Species (Pergamon, Ox-
ford, 1982).
34. D. Rabinowitz, S. Cairns, T. Dillon, in Conservation Biology, M. E. Soule, Ed.
(Sinauer, Sunderland, MA, 1986), pp. 182-204.
35. The 177 plant species in the British Flora are clearly unlikely to be a random subset
of i:he entire 1822 plant species in Britain. Understandably, very widespread species
tend to be overrepresented, at the expense of middling species. Species with highly
restricted distributions, however, form about 30% both of the British Flora and of
the entire flora. For further details about possible biases, and about the consensual
process among 15 ecologists and systematists, see (34).
36. The category of "rare habitat'' may be overrepresented in the British Flora by virtue
of the country's long coastline and many maritime species.
37. See, fur example, the discussion of sparse and patchily distributed species in species-
rich plant communities by P. J, Grubb [in Community Ecology, J. M. Diamond and
T. J. Case, Eds. (Harper and Row, New York, 1986), pp. 207-226] and of
experimental design in demographic studies of rare plants by J. Travis and R. Sutter
[Natural Areas]. 6, 3 (1987)].
38. V. Pearse, Living Invertebrates (Blackwell, Oxford, 1987).
39. There are, of course, varying opinions about what constitutes a fundamentally
diJferent body plan, which is why different schemes recognize from 25 to 35 animal
phyla.
40. G. E. Hutchinson, Am. Nat. 93, 145 (1959).
41. These ideas were developed further by E. C. Pielou, Biogeography (Wiley, New
York, 1979).
42. G. C. Ray, Am. Zoo/. 25, 451 (1985).
43. Notable examples are provided by Erophila and Arabidopsis, British plants in the
mustard family.
44. R. K. Selander, in Population Genetics and Molecular Evolution, T. Ohta and K. Aoki,
Eds. (Springer-Verlag, Berlin, 1985), pp. 85-106.
45. J. M. Diamond, Nature 315, 538 (1985).
The Ecological Context of
Life History Evolution
LINDA PARTRIDGE AND PAUL
Life histories are the probabilities of survival and the rates
of reproduction at each age in the life-span. Reproduction
is costly, so that fertility at all ages cannot simultaneously
be maximized by natural selectio11. Allocation of repro-
ductive effort has evolved in response to the demographic
impact of different environments but is constrained by
genetic variance and evolutionary history.
E
VEN THE MOST FAMILIAR ORGANISMS HAVE VERY DIVERSE
life histories. Most small. birds, such as chickadees or great
tits, breed in the spring following their birth, and continue
to nest every year until their death. As adults, they have a 50 percent
chance of surviving each successive winter. In sharp contrast, most
Pacific salmon breed in a suicidal burst as 3-year-olds. Oak trees
have high adult survival rates, take more than 3 years before
producing even their first few acorns, but then step up production
imtil their acorns are numbered in thousands each year.
Making such diversity intelligible is one reason for studying life
history evolution. Another is to predict the ways in which popula-
tions will respond to changed environments, including harvesting.
Understanding life history diversity means facing fundamental
16 SEPTEMBER 1988
46. P. H. Raven, Bull. Entomol. Soc. Am. 29, 5 (1983).
47. C. A. Toft, in Community Ecology, J, M. Diamond and T. J. Case, Eds. (Harper and
Row, New York, 1986), pp. 445-463. .
48. In an essentially one-dimensional enviromnent with ftactal dimension D, Hutchin-
son and MacArthur's (20) argument suggests a scalin~f S - L -D; in an essentially
two-dimensional enviromnent, the scaling is S - L- . When combined with the
estimateD- 1.5 (21), this gives the result in the text.
49. T. L. Erwin andJ. C. Scott, Co/eopt. Bull. 34,305 (1980); T. L. Erwin, ibid. 36,74
(1982); Bull. Entomol. Soc. Am. 29, 14 (1983).
50. R. Howard, personal communication, cited in (43).
51. See, for example, N. E. Stork, Ecol. Entomol. 12, 69 (1987).
52. I suggest 1 haas the smallest meaningful area for such a study, although not the
least of our problems is that we do not understand just what the smallest scale need
be to give sensible answers.
53. A. P. Dobson, personal communication.
54. Table 3 was compiled by A. P. Dobson, with help from A. M. Lyles and A.
Merenlender. The number of species is taken from R. F. K. Barnes, A Synoptic
Classification of Living Organisms (Blackwell, Oxford, 1983); most of the numbers
are fur recorded species, but some (especially nematodes and insects, as emphasized
by the question mark) are Barnes's estimates. The numbers of publications come
from The Zoological Record, averaged over the years 1978 through 1987 (with the
coefficient of variation of the annual number shown in parentheses). The Zoological
Record lists papers covering every aspect of the zoology of animals (scanning over
6000 journals and other documents), except for insects, where only publications
dealing with systematics, taxonomy, or nomenclature are given; Table 3 thetefure
underestimates, by an unknown factor, work on insects relative to that on other
animals.
55. This article has been shaped by helpful suggestions from R. K. Colwell, P. J.
DeVries, W. D. Hamilton, A. P. Dobson, H. S. Hom, M. Kreitman, J. H.
Lawton, J. May, C. A. Toft, and two anonymous reviewers. Supported in part by
NSF grant DMS87-03503.
H. HARVEY
questions about the functioning of organisms: What determines the
maximum possible rate of reproduction? What developmental and
physiological processes would have to be altered to increase the
potential life-span? Why is rapid growth during development often
associated with an elevated risk of morality? Answers to these kinds
of questions are important in agricultural production and medicine,
as well as ecology. Nevertheless, the subject is firmly rooted in
ecology because, as we shall see, life histories evolve largely in
response to the impact of different environments on the survival and
fertility of different age-classes.
Our aim in this article is to evaluate the successes and limitations
of the adaptationist approach to understanding life history evolu-
tion. It has been claimed that such an approach is doomed (1). In
contrast, we shall argue that, when appropriately handled, it can
have considerable utility for understanding both the diversity of life
histories and the mechanisms constraining their form. We do not
provide a comprehensive account, for which reviews are already
available (2-4). We first outline the demographic model underlying
most adaptationist interpretations of life history variation before
going on to show how optimal life histories might be realized. We
L. Partridge is a reader in the Department of Zoology, University of Edinburgh, West
Mains Road, Edinburgh EH9 3JT, UK. P. H. Harvey is a lecturer in the Department
of :ZOology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.
ARTICLES 1449